The Possible Pollen Cone of the Late Triassic Conifer Heidiphyllum/Telemachus (Voltziales) from Antarctica

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The Possible Pollen Cone of the

by Benjamin Bomfleur, Rudolph Serbet, Edith L. Taylor, and Thomas N. Taylor

2011

This is the published version of the article, made available with the permission of the publisher. The original published version can be found at the link below.

Bomfleur, B., Serbet, R., Taylor, E., and Taylor, N. 2011. The Possible Pollen Cone of the Late Triassic Conifer Heidiphyllum/Telemachus (Voltziales) From Antarctica. Antarctic Science 23(4): 379-385.

Published version: http://dx.doi.org/10.1017/S0954102011000241

This work has been made available by the University of Kansas Libraries’ Office of Scholarly Communication and Copyright. Antarctic Science 23(4), 379–385 (2011) & Antarctic Science Ltd 2011 doi:10.1017/S0954102011000241 The possible pollen cone of the Late Triassic conifer Heidiphyllum/Telemachus (Voltziales) from Antarctica BENJAMIN BOMFLEUR, RUDOLPH SERBET, EDITH L. TAYLOR and THOMAS N. TAYLOR Division of Paleobotany at the Department of Ecology and Evolutionary Biology, and Natural History Museum and Biodiversity Institute, University of Kansas, Lawrence, KS 66045, USA bbomﬂ[email protected]

Abstract: Fossil leaves of the Voltziales, an ancestral group of conifers, rank among the most common plant fossils in the Triassic of Gondwana. Even though the foliage taxon Heidiphyllum has been known for more than 150 years, our knowledge of the reproductive organs of these conifers still remains very incomplete. Seed cones assigned to Telemachus have become increasingly well understood in recent decades, but the pollen cones belonging to these Mesozoic conifers are rare. In this contribution we describe the ﬁrst compression material of a voltzialean pollen cone from Upper Triassic strata of the Transantarctic Mountains. The cone can be assigned to Switzianthus Anderson & Anderson, a genus that was previously assumed to belong to an enigmatic group of pteridosperms from the Triassic Molteno Formation of South Africa. The similarities of cuticle and pollen morphology, together with co-occurrence at all known localities, indicate that Switzianthus most probably represents the pollen organ of the ubiquitous Heidiphyllum/Telemachus plant.

Received 9 December 2010, accepted 23 February 2011, ﬁrst published online 5 April 2011

Key words: coniferophytes, Gondwana, in situ pollen, Switzianthus, systematic palaeobotany, Transantarctic Mountains

Introduction (Anderson 1978, Retallack 1981, Yao et al. 1993, Axsmith et al. 1998, Nielsen 2005, Escapa et al.2010).Further The Voltziales encompasses a diverse array of extinct evidence for Triassic transitional conifers from the Southern conifers that are believed to be transitional between the Hemisphere has come from permineralized deposits of Palaeozoic conifers and the modern Coniferales (e.g. Florin the central Transantarctic Mountains, which have yielded 1938–45, Rothwell et al. 2005, Taylor et al. 2009). The various anatomically preserved conifer organs, including majority of voltzialean taxa have been described from the stems and leaves of Notophytum krauselii Meyer-Berthaud upper Palaeozoic (e.g. Rothwell et al. 2005). There are, & Taylor (Meyer-Berthaud & Taylor 1991, Axsmith et al. however, fossils of this ancient conifer group that occur in 1998), seed cones of Parasciadopitys aequata Yao, Taylor deposits as young as the Early Jurassic (Taylor et al. 2009), & Taylor (Yao et al. 1997), and the pollen cone Leastrobus or possibly even Early Cretaceous (Miller 1977). fallae Hermsen, Taylor & Taylor (Hermsen et al. 2007). The Several Northern Hemisphere representatives of Mesozoic affinities and interrelationships of these fossils continue to voltzialean conifers have been intensely studied and are remain largely obscure. At least some of these morphotaxa, known in remarkable detail (e.g. Grauvogel-Stamm 1978). however, are regarded as being the permineralized By contrast, knowledge about the Gondwanan Voltziales has equivalents of the compression taxa Heidiphyllum and remained very incomplete. This is remarkable, because Telemachus (e.g. Axsmith et al. 1998, Escapa et al.2010, voltzialean conifer leaves have been described from the Schwendemann et al.2010). Gondwanan Triassic since the first half of the 19th century Even though voltzialean foliage and seed cones are (e.g. Morris 1845, Feistmantel 1889). These leaves have common elements in Gondwanan Triassic floras and have initially been assigned to a variety of morphogenera, (in part) been known for more than 150 years, information including Zeugophyllites Brongniart, Podozamites Braun, on the corresponding pollen cones has remained very Phoenicopsis Heer, Desmiophyllum Lesquereux, and limited. This may be due to the small size and ephemeral Noeggerathiopsis Feistmantel, but the vast majority were nature of these pollen cones (Rothwell & Mapes 2001, later merged into a single species Heidiphyllum elongatum Hermsen et al. 2007). In this contribution we present the (Morris) Retallack (Retallack 1981). Heidiphyllum is first record of voltzialean pollen cone compressions from currently recognized as the most widespread conifer leaf the Triassic of Antarctica. The specimens are assigned to in the Triassic of the Southern Hemisphere. It is frequently Switzianthus Anderson & Anderson, a morphogenus that found together with the supposedly affiliated seed was previously thought to belong to an enigmatic group of cone Telemachus Escapa, Decombeix, Taylor & Taylor pteridosperms from the Molteno Formation of South Africa

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Fig. 2. Voltzialean foliage and seed cone from the Upper Triassic of the Allan Hills, south Victoria Land, East Antarctica. Scale bars 5 1 cm. a. Heidiphyllum elongatum (Morris) Retallack. T11-582a. b. Telemachus antarcticus Escapa, Decombeix, Taylor & Taylor. T11-411. Fig. 1. Locality map showing the fossil sites in south Victoria Land and the central Transantarctic Mountains, East Antarctica. Light grey tones indicate areas of rock outcrop. (Level 14, Profile 2 in Barrett 1969) with intercalated coaly layers and thin sheets of light-grey, very fine-grained (Anderson & Anderson 2003). By contrast, the similarities sandstone. The Falla Formation is interpreted as deposits in cuticle and pollen morphology, together with the co- of a low-sinuosity braided-stream system with a high occurrence at all known localities, lead us to the conclusion proportion of lacustrine and paludal overbank fines (e.g. that Switzianthus probably represents the pollen cone of the Barrett et al. 1986). A single additional specimen comes ubiquitous Heidiphyllum/Telemachus plant. from Member C of the Lashly Formation in the Allan Hills, south Victoria Land (Fig. 1). At both sites, the specimens co-occur with abundant Heidiphyllum leaves (Fig. 2a) Material and methods and Telemachus seed cones (Fig. 2b). The age of the Most specimens were collected from the type section plant-bearing horizons is considered to be Carnian or of the Falla Formation on the northern flank of the western Norian (early or middle Late Triassic) based on microfloras ridge of Mount Falla (Queen Alexandra Range, central (Kyle & Schopf 1982, Farabee et al. 1989). Transantarctic Mountains; 84822'S, 164855'E) (Fig. 1). Hand specimens were photographed with a Nikon D100 Plant fossils occur within a 16 m thick mudstone interval digital camera. Macroimages were taken with a Leica DC500

Fig. 3. Gross morphology, cuticle features, and in situ pollen of the voltzialean pollen cone Switzianthus sp. from the Upper Triassic of the Transantarctic Mountains. a. Intact pollen cone. Note that the spiny aspect of some sporophyll apices along the cone margin is due to individual superpositioned sporophylls being cross-sectioned. Mount Falla, T7-185a. Scale bar 5 5 mm. b. Counterpart of the same specimen. T7-185b. Scale bar 5 5 mm. c. Partially disintegrated pollen cone. Mount Falla, T7-172b. Scale bar 5 5 mm. d. Comparatively intact pollen cone. Allan Hills, T8-229b. Scale bar 5 5 mm. e&f.Details of individual ovate distal microsporophyll laminae with a slightly displaced central pedicel scar, a surrounding rugose area, and a distal portion with ﬁne radiating striations. Scale bar 5 1 mm. g. Cuticle of the presumably outer (i.e. abaxial) surface of the distal microsporophyll lamina from a partially disintegrated pollen cone from Mount Falla (T5-111), showing the outlines of more or less longitudinally aligned, elongated rectangular to polygonal cells. Slide #24033. Scale bar 5 50 mm. h. Large cluster of Alisporites Daugherty type pollen grains isolated from a partially disintegrated cone from Mount Falla (T5-111). Slide #24034. Scale bar 5 100 mm. i–k. Individual pollen grains in polar (i & j) and lateral (k) view. Slides #24033 (j & k) and #24037 (i). Scale bars 5 25 mm.

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http://journals.cambridge.org Downloaded: 19 May 2014 IP address: 129.237.46.100 382 BENJAMIN BOMFLEUR et al. digital camera mounted on a Leica MZ16 stereo dissecting the entire grain; cappa generally thickened, with smooth or microscope. Cuticle and in situ pollen samples were freed finely reticulated to rugose surface; distal surface with a from the matrix using 48% hydrofluoric acid (HF) for several sharply defined, narrow-rectangular longitudinal sulcus of days, and then carefully neutralized by repeated washing with c. 10–12 mm width (Fig. 3i & j). distilled water. Organic residues were further macerated Remarks: The material agrees in all comparable features using Schulze’s reagent (40% nitric acid (HNO )withafew 3 with the pollen cone genus Switzianthus from the Upper crystals of potassium chlorate (KClO )) for several hours, 3 Triassic Molteno Formation of South Africa, including: and subsequently cleaned and bleached using a 4% potassium 1) organization of a more or less compact cone, 2) the size hydroxide solution (KOH ) for a few seconds. Cuticle and aq range (2.8–4.5 cm long), 3) number of scales per helix, pollen were finally dehydrated in glycerol and mounted on 4) dimension, morphology and texture of the sporophylls, permanent slides using glycerol jelly. and 5) the in situ pollen grains. Compared to the South Hand specimens and slides are stored in the Paleobotanical African specimens, the cuticle fragments probably derive Collections of the Department of Ecology and Evolutionary from the distal microsporophyll portion (see Anderson & Biology, and Natural History Museum and Biodiversity Anderson 2003 pl. 56, fig. 1). Institute, University of Kansas, Lawrence (KS), USA. Two species have been described in the South African material, i.e. S. moriformis Anderson & Anderson and Systematics S. crispiformis Anderson & Anderson. The present material Phylum CONIFEROPHYTA more closely resembles S. crispiformis due to its relatively Class CONIFEROPSIDA small size and well-defined pedicel scar. However, as both Order VOLTZIALES species lack well-defined diagnostic criteria and remain Family incertae sedis poorly understood, we refrain from attempting a species Genus Switzianthus Anderson & Anderson determination until additional and better preserved material Type species. Switzianthus moriformis Anderson & becomes available. Anderson, 2003 Essential details on pollen-sac morphology and attachment in Switzianthus remain uncertain. Following Anderson & Switzianthus sp. Anderson (2003), we suggest that the pitted area around the (Fig. 3a–k) pedicel scar on the adaxial microsporophyll surface may be the Material: Eight specimens from Mount Falla on the area of former pollen-sac attachment. In none of the specimens following seven slabs: T5-69, 79, 86, 111, T7-161, 172, so far described, however, are the actual pollen sacs preserved. 185 (Upper Triassic, Falla Formation, Mount Falla, Queen This may indicate that the pollen-sac material was very delicate Alexandra Range; 84822'S, 164855'E); one additional and already withered away when the cones became buried. specimen T8-229 from the Allan Hills (Upper Triassic, In situ palynological samples were taken by carefully Member C, Lashly Formation, Allan Hills, south Victoria removing organic material from the microsporophylls of a Land). Cuticle and pollen preparations of specimen T5-111 disintegrated cone. The recovered palynomorphs consist on slides #24026–24038. exclusively of microsporophyll cuticle fragments and the above described bisaccate pollen grains, many of the latter Description: Cylindrical pollen cones, 2.7–c. 4 cm long and occurring in distinct clusters (Fig. 3h). 0.8–1.5 cm in diameter, more or less compact, composed of helically arranged microsporophylls with each helix comprising c. 6–10 scales (Fig. 3a–d). Scales robust, Discussion peltate, each with a delicate pedicel and an ovate distal Switzianthus has previously been affiliated with the portion with an obtuse to acuminate apex (Fig. 3a–c, e & f); enigmatic and poorly defined pteridosperm leaf Dejerseya distal microsporophyll portions c. 4–6 mm long and 3–5 mm Herbst, based on what have been described as similar wide; adaxial surface with a circular pedicel scar of c.1mm cuticle features and co-occurrence at some localities. By diameter positioned slightly below the centre, a surrounding contrast, we interpret Switzianthus as a voltzialean pollen thickened area with a granulose surface, and a thin, woody cone that probably belongs to the Antarctic Heidiphyllum/ distal portion with radiating striations (Fig. 3e & f). Cuticle Telemachus conifer. This determination is based on: fragments thin, showing a pattern of more or less 1) structural correspondence to other voltzialean pollen longitudinally aligned, elongated rectangular to polygonal cones, 2) similarities in cuticle and pollen morphology, and cells 30–40 mm long and 20–30 mm wide (Fig. 3g). 3) co-occurrence at all known localities. Pollen grains bisaccate, measuring between 60 x 38 mm and 100 x 60 mm (Fig. 3h–k). Sacci narrow-elliptical in Structural correspondence polar view (Fig. 3h–j), proximally attached in equatorial region, distally inclined (Fig. 3k). Corpus outline circular to The morphology of Switzianthus is unlike that of any narrow-elliptic in polar view, approximately half as wide as known pteridosperm pollen organ. By contrast, the helical

http://journals.cambridge.org Downloaded: 19 May 2014 IP address: 129.237.46.100 A TRIASSIC POLLEN CONE FROM ANTARCTICA 383 arrangement of peltate microsporophylls with ovate, are hypostomatic, and the stomata are surrounded by a bilaterally symmetrical distal portions indicates affinities ring of five to six strongly cutinized subsidiary cells. with either conifers or cycads (e.g. Taylor et al.2009). Guard cell cutinizations are not preserved. The cuticle of A cycadalean affinity of Switzianthus can be excluded regular epidermal cells and subsidiary cells has been because fossil cycads are known to have produced simple, described to be nonpapillate. Judging from the illustrations, monocolpate Cycadopites Wodehouse pollen, whereas however, papillae occur on the lower sporophyll surface of in situ pollen of Switzianthus is bisaccate non-taeniate, and at least some of the specimens (Anderson & Anderson 2003 of the Alisporites Daugherty type (5 Alisporites sensu lato; pl. 56, fig. 1). see Balme 1995, Anderson & Anderson 2003). Although these features agree to some extent with the Structurally, Switzianthus agrees with many of the well- cuticle morphology of Dejerseya (Anderson & Anderson known Triassic pollen-cone compressions from the Northern 1989), as suggested by Anderson & Anderson (2003), they Hemisphere, including Ruehleostachys Roselt emend. Arndt are also well in accordance with those of Heidiphyllum and (5 Willsiostrobus Grauvogel-Stamm & Grauvogel), Darneya Telemachus. Heidiphyllum leaves are hypostomatic to weakly Schaarschmidt & Maubeuge emend. Grauvogel-Stamm, and amphistomatic (Anderson 1978, Anderson & Anderson 1989, Sertostrobus Grauvogel-Stamm (see Grauvogel-Stamm 1978, Axsmith et al. 1998). In Heidiphyllum and Telemachus,the Arndt 2002). These taxa are distinguished primarily by the stomata are surrounded by a ring of five to six variably morphology and mode of attachment of the pollen sacs thickened subsidiary cells (Anderson 1978, Anderson & (e.g. Grauvogel-Stamm & Schaarschmidt 1979). Because Anderson 1989, Yao et al. 1993, Axsmith et al.1998).The evidence for pollen-sac morphology and attachment in development of papillae on regular epidermal cells and Switzianthus is still lacking, more detailed comparisons subsidiary cells is variable in Heidiphyllum, with papillae on cannot be made. Several types of voltzialean pollen cones the lower leaf surface being strongly developed (e.g. Axsmith have recently also been described from the Molteno et al. 1998), weakly developed (e.g. Anderson 1978 pl. 9, Formation of South Africa (Anderson & Anderson 2003). fig. 5), or absent (Anderson & Anderson 1989 pl. 251, figs 6 These differ from Switzianthus primarily in the larger size & 7). Papillae are also absent in Telemachus (Yao et al. and the more complex cone structure, some with up to 30 1993). Hence, the cuticle features of Switzianthus,which variably ornamented microsporophyll scales per helix were suggested as indicating affinities with the pteridosperm (Anderson & Anderson 2003). Dejerseya, are at least as close to those of the Voltziales. Until now only two conifer pollen cones are known from The bisaccate, non-taeniate, sulcate pollen of Switzianthus the Triassic of Antarctica. The voltzialean pollen cone corresponds to the dispersed pollen genus Alisporites (e.g. Leastrobus fallae is based on anatomically preserved Balme 1995). Yao et al. (1993) reported large numbers of material from an outcrop at the base of Mount Falla similar grains attached to the cuticles of Telemachus cones, (Hermsen et al. 2007). The cone structure of Switzianthus which led these authors to suggest that the parent plant of corresponds well with that of Leastrobus, although the Telemachus may have produced Alisporites-type pollen (Yao latter appears to be much smaller, with microsporophyll et al. 1993). This interpretation is further supported by the fact laminae measuring only up to 1.2 mm wide (Hermsen et al. that several voltzialean pollen cones from the Triassic of the 2007). Cantrill et al. (1995) described charcoalified remains Northern Hemisphere, e.g. Ruehleostachys bromsgrovensis of an undetermined conifer cone from the Upper Triassic of (Grauvogel-Stamm) Roselt emend. Arndt, R. rhomboidalis the Amery Group, Prince Charles Mountains. This cone is (Grauvogel-Stamm) Roselt emend. Arndt, and R. willsii characterized by helically arranged, peltate scales with (Townrow) Roselt emend. Arndt, produced Alisporites-type obovate distal portions that are superficially similar to those pollen as well (see Grauvogel-Stamm 1978, Balme 1995, of Switzianthus. The size range of the microsporophylls, Arndt 2002). Also the Antarctic cone Leastrobus fallae with distal portions being 0.6–3.2 mm long by 0.5–2.3 mm contains morphologically similar pollen. The grains differ wide, appears to be somewhat intermediate between those mainly in the smaller size and in the more clearly developed, of Leastrobus and the specimens described here. Hence, fine corpus reticulation (Hermsen et al. 2007). these three Antarctic taxa may be interpreted as forming an intergrading series within a group of structurally similar Co-occurrence conifer pollen cones. Switzianthus has been affiliated with Dejerseya foliage in part because of the co-occurrence at some localities in Similarities in cuticle and pollen morphology South Africa (Anderson & Anderson 2003). A particular The cuticles recovered from the Antarctic cones are poorly reference for this hypothesis is supposed to be a large preserved and demonstrate only few diagnostically relevant collection of plant fossils from the Lit111 site, where details. Well-preserved cuticles of microsporophylls of Dejerseya leaves account for c. 20% of the foliage taxa and Switzianthus have been described from South Africa Switzianthus is the most common pollen organ (Anderson (Anderson & Anderson 2003). The sporophyll laminae & Anderson 2003). However, nineteen other gymnosperm

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Table I. Comparison of localities where Switzianthus Anderson & Anderson has been found (data for South Africa are taken from Anderson & Anderson 2003). Region Locality Sample size Dejerseya Herbst Heidiphyllum Retallack (no. of slabs) Antarctica Mount Falla, Level 14 , 150 11 Allan Hills , 900 21 South Africa Little Switzerland, 111 2173 11 Aasvoe¨lberg, 411 2176 21 Matatiele, 111 1082 21 Winnaarspruit, 111 64 11

foliage genera and not less than six other pollen organ valuable discussion, and gratefully acknowledge helpful genera occur at the Lit111 site. An affiliation based on the comments and suggestions by Brian Axsmith (Mobile, quantitative occurrences of individual taxa in this assemblage AL, USA) and David Cantrill (Melbourne, Australia). must therefore remain questionable. Moreover, Switzianthus Financial support has been provided by the Deutsche is now known from several extensively sampled plant fossil Forschungsgemeinschaft (DFG grant KE584/16-2) and in localities in which Dejerseya is absent (Table I). This is part by the National Science Foundation (NSF grant DPP remarkable, as it can be assumed that the pollen organs of an 88-15976 to E.L.T. and T.N.T). individual plant are far less likely to become preserved in a fossil assemblage than the affiliated foliage (e.g. Rothwell & Mapes 2001). References By contrast, Switzianthus is associated with Heidiphyllum ANDERSON, H.M. 1978. Podozamites and associated cones and scales from leaves at all known localities (Table I). In the present the Upper Triassic Molteno Formation, Karoo Basin, South Africa. collections from Allan Hills and Mount Falla, Heidiphyllum Palaeontologia Africana, 21, 57–77. forms a dominant component on many of the slabs. The ANDERSON, J.M. & ANDERSON, H.M. 1989. Palaeoflora of southern Africa: intact Switzianthus cone from Allan Hills occurs on a slab Molteno Formation (Triassic). Vol. 2. Gymnosperms (excluding Dicroidium). Rotterdam: Balkema, 567 pp. that otherwise bears only Heidiphyllum leaves. ANDERSON, J.M. & ANDERSON, H.M. 2003. Heyday of the gymnosperms: systematics and biodiversity of the Late Triassic Molteno fructifications. Pretoria: National Botanical Institute, Strelitzia 15, 399 pp. Conclusions ARNDT, S. 2002. Morphologie und Systematik ausgewa¨hlter Mesozoischer Taken together, the evidence leads us to the conclusion Koniferen. Palaeontographica, B262, 1–23. AXSMITH, B.J., TAYLOR, T.N. & TAYLOR, E.L. 1998. Anatomically preserved that Switzianthus is a voltzialean pollen cone that probably leaves of the conifer Notophytum krauselii (Podocarpaceae) from the belonged to the same parent plants that produced the Triassic of Antarctica. American Journal of Botany, 85, 704–713. Heidiphyllum leaves and Telemachus seed cones of the BALME, B.E. 1995. Fossil in situ spores and pollen grains: an annotated Antarctic Triassic. Only a single morphospecies of catalogue. Review of Palaeobotany and Palynology, 87, 81–323. Heidiphyllum, i.e. H. elongatum, is currently recognized BARRETT, P.J. 1969. Stratigraphy and petrology of the mainly fluviatile Permian and Triassic Beacon rocks, Beardmore Glacier area, Antarctica. in the Triassic of Antarctica and South Africa (Anderson & Columbus, OH: Ohio State University Research Foundation, Institute of Anderson 2003). At the same time, however, at least five Polar Studies, Report No. 34, 132 pp. different species of voltzialean pollen cones have been BARRETT, P.J., ELLIOT, D.H. & LINDSAY, J.F. 1986. The Beacon Supergroup described from the Molteno Formation, and possibly two (Devonian–Triassic) and Ferrar Group (Jurassic) in the Beardmore further distinct taxa are known from the Triassic of Glacier area, Antarctica. Antarctic Research Series, 36, 339–428. CANTRILL, D.J., DRINNAN, A.N. & WEBB, J.A. 1995. Late Triassic plant Antarctica (Cantrill et al. 1995, Hermsen et al. 2007). If fossils from the Prince Charles Mountains, East Antarctica. Antarctic future studies should confirm that our interpretation of Science, 7, 51–62. Switzianthus as yet another type of pollen cone to be ESCAPA, I.H., DECOMBEIX, A.-L., TAYLOR, E.L. & TAYLOR, T.N. 2010. affiliated with H. elongatum is accurate, this would indicate Evolution and relationships of the conifer seed cone Telemachus: that a remarkable, yet previously unrecognized diversity evidence from the Triassic of Antarctica. International Journal of Plant Sciences, 171, 560–573. and morphological disparity of natural conifer species may FARABEE, M.J., TAYLOR, T.N. & TAYLOR, E.L. 1989. Pollen and spore be hidden among the uniform morphology of voltzialean assemblages from the Falla Formation (Upper Triassic), central foliage and seed cones in the Triassic of Gondwana. Transantarctic Mountains, Antarctica. Review of Palaeobotany and Palynology, 61, 101–138. FEISTMANTEL, O. 1889. U¨ bersichtliche Darstellung der geologisch- Acknowledgements palaeontologischen Verha¨ltnisse Su¨d-Afrikas. I. Theil. Die Karoo- Formation und die dieselbe unterlagernden Schichten. Abhandlungen We wish to thank Ignacio Escapa (Trelew, Chubut, der Ko¨niglichen Bo¨hmischen Gesellschaft der Wissenschaften Argentina) and Hans Kerp (Mu¨nster, Germany) for Mathematisch-Naturwissenschaftliche Klasse VII, 3, 1–89.

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